Aposematism and the handicap principle.

Aposematic prey use conspicuous warning signals to advertise their defenses to predators. It has long been recognized that the efficiency of a warning signal may be reduced if poorly defended prey (automimics) are present in the population. The handicap principle suggests that the use of warning signals by poorly defended prey may be kept in check if signaling is costly. Three mechanisms that involve signal costs have been proposed to facilitate honest warning signals in prey: go-slow behavior in predators, resource allocation trade-offs, and costs of detection alone. We study all three in a unified game-theoretical framework. We find that the go-slow mechanism and the resource allocation mechanism can introduce differential benefits and differential costs of signaling, respectively, and can support honest signaling in accordance with the handicap principle. When honest signaling is maintained by the go-slow mechanism, conspicuous prey will necessarily suffer more attacks on average than cryptic prey. In contrast, when honest signaling is maintained by the resource allocation mechanism, cryptic prey will suffer more attacks. The detection cost mechanism lacks differential costs and benefits, and its potential for maintaining honest signaling equilibria is limited. We relate our results to intra- and interspecific correlations between conspicuousness and defense.